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CHAPTER 9: FUNGAL GENETICS,
MOLECULAR GENETICS AND GENOMICS
Chapter 9 is divided into the
following major sections:
overview: the place
of fungi in genetical research
Neurospora and
classical (Mendelian) genetics
structure and organisation of the
fungal genome
genetic variation in fungi
applied molecular genetics of fungi
Returning to the genome
expressed sequence tags and
microarray technology
SAMPLE
TEXT:
In this
chapter we cover the basic and applied genetics of fungi,
including the features that continue to make fungi
important model organisms for genetical research. The
chapter includes recent molecular approaches in a range
of fields such as the analysis of fungal pathogenicity
determinants and the development of fungi as
factories for foreign gene products. It also
covers the roles of extrachromosomal genes in
ageing-related senescence and the effects of fungal
viruses (hypoviruses) in suppressing pathogenic
virulence.
Fungi
are eminently suitable for biochemical studies because of
their simple nutrient requirements, and because
classical genetics has provided excellent
physical maps of the chromosomal genes. Studies on one
fungus in particular Neurospora crassa
led to the classical concept of one gene,
one enzyme, for which Beadle & Tatum (1945)
received the Nobel Prize. However, with more recent
knowledge it is more accurate to say that one gene
can encode one enzyme the situation is
complicated because gene splicing occurs to remove
non-coding introns in the pre-messenger RNA.
At
the time of writing (May 2004), the genomes of nearly 150
organisms have been sequenced mainly bacteria and
archaea, but also the genomes of mouse and
man. Notably in our present context, the
high-quality draft genome sequences
of five fungi have been published Saccharomyces
cerevisiae, Neurospora crassa, Emericella
(Aspergillus) nidulans, Schizosaccharomyces
pombe and the rice blast pathogen, Magnaporthe
grisea. The first four of these are Ascomycota with
well-mapped chromosomes, providing a basis for combining
classical and molecular genetics.
Chapter
9 images. Click on thumbnails
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Fig. 9.1
Fig. 9.8
Fig. 9.10
Fig. 9.14
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Fig. 9.2
Fig. 9.5
Fig. 9.11
Fig. 9.15
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Fig. 9.3
Fig. 9.6
Fig. 9.12
Fig. 9.16
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Fig. 9.4
Fig.
9.7
Fig. 9.9
Fig. 9.13
[see text
for
Figs 9.17 and 9.18]
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Table
9.1. Reported chromosome counts in some
representative fungi |
Oomycota |
|
Phytophthora
spp. (many) |
9-10 |
Achlya
spp. |
3, 6, 8 |
Saprolegnia spp. |
8-12 |
Pythium |
commonly 10 or 20 |
Chytridiomycota |
|
Allomyces
arbuscula |
16 |
A. javanicus |
14 (variable in
hybrids and polyploids) |
Ascomycota |
|
Schizosaccharomyces
pombe |
3 |
Neurospora
crassa |
7 |
Saccharomyces
cerevisiae |
16 |
Emericella (Aspergillus)
nidulans |
8 |
Coccidioides
posadasii |
4 |
Trichophyton
rubrum |
4 |
Magnaporthe
grisea |
7 |
Basidiomycota |
|
Filobasidiella
neoformans |
11 |
Schizophyllum
commune |
11 |
Coprinus
cinereus |
13 |
Puccinia
kraussianna |
30-40 |
--------------------------------------------------
Table
9.2. Some reported (approximate) genome sizes of
fungi and fungus-like organisms |
Aspergillus
fumigatus (potential human pathogen) |
30 Mb |
A. niger
(industrially important: citric acid, enzyme
production) |
30 Mb |
Candida
albicans (human commensal and potential
pathogen) |
16 Mb |
Filobasidiella
(Cryptococcus) neoformans (human
pathogen) |
21 Mb |
Emericella (Aspergillus)
nidulans (experimental model fungus) |
28 Mb |
Neurospora
crassa (experimental model fungus) |
40 Mb |
Phanaerochaete
chrysosporium (wood-decay Basidiomycota) |
40 Mb |
Phytophthora
infestans (plant pathogen; Oomycota) |
240 Mb |
Phytophthora
sojae (pathogen of soybean; Oomycota) |
62 Mb |
Pneumocystis
jiroveci (pathogen of immunocompromised
humans) |
7.7 Mb |
Saccharomyces
cerevisiae (brewing and breadmaking yeast) |
12 Mb |
Schizosaccharomyces
pombe (experimental model; fission yeast) |
14 Mb |
-----------------------
Table
9.4. A list of fungi initially proposed to form
the basis of a co-ordinated genome sequencing
effort in the USA |
Organisms bycategory |
Significance (and
chapter reference)
[See footnotes to this table] |
Estimated genome (Mb) |
MEDICINE |
|
Filobasidiella
(Cryptococcus) neoformans serotype A |
Basidiomycota. Encapsulated
yeast; causes fatal meningitis in humans |
24 Mb on 11 chromosomes |
Coccidioides posadasii |
Ascomycota. Soil fungus
endemic to southwestern USA; causes fatal human
infection; also a bioterrorism threat |
29 Mb on 4 chromosomes |
Pneumocystis carinii (human
and mouse forms) |
The leading opportunistic
pathogen of AIDS patients; drug resistance is
emerging |
7.5; 6.5 Mb |
Trichophyton rubrum |
Ascomycota. The most common
fungal infection in the world; adapted for growth
on human skin |
12 Mb on 4 chromosomes |
Rhizopus oryzae |
Zygomycota. Can cause
infection of humans (zygomycosis) |
36 Mb |
COMMERCE |
|
Magnaporthe grisea |
Causes rice blast disease.
A model fungal plant pathogen |
40 Mb on 7 chromosomes |
Aspergillus flavus |
Ascomycota/ mitosporic
fungus. Source of aflatoxin and one cause of
human aspergillosis |
40 Mb on 8 chromosomes |
Emericella (Aspergillus)
nidulans |
Ascomycota. Key model
system for genetics and cell biology. (Already
part-sequenced) |
31 Mb on 8 chromosomes |
Aspergillus terreus |
Mitosporic fungus. Major
source of the cholesterol-lowering drug,
lovastatin |
30 Mb |
Fusarium graminearum |
Mitosporic fungus. Causes
head blight on wheat and barley; produces
mycotoxins. |
40 Mb on 9 chromosomes |
EVOLUTION/ FUNGAL
DIVERSITY |
|
Neurospora discreta |
Ascomycota. Fungal model
for population genetics and comparison with N.
crassa (already sequenced) |
40 Mb on 7 chromosomes |
Coprinus cinereus |
Basidiomycota. Model for
fungal differentiation produces toadstools
|
37.5 Mb on 13 chromosomes |
Batrachochytridium
dendrobatidis |
Chytridiomycota. Recently
described fungus that causes widespread
population decline of amphibians |
30 Mb on 20 (?) chromosomes |
Ustilago maydis |
Basidiomycota. Model for
plant-pathogen interactions |
20 Mb |
Paxillus involutus |
Basidiomycota. Symbiotic
mycorrhizal fungus of many trees, easily
manipulated in laboratory conditions |
40 Mb |
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